Geol. Mag. 140 (5 ), 2003, pp. 523–538. c© 2003 Cambridge University Press 523DOI: 10.1017/S0016756803007854 Printed in the United Kingdom

Palaeobiogeographic implications of Middle Permian brachiopodsfrom Johore (Peninsular Malaysia)

MASATOSHI SONE*†, IAN METCALFE* & MOHD SHAFEEA LEMAN*‡*Asia Centre, University of New England, Armidale, NSW 2351, Australia

‡School of Environmental Sciences and Natural Resources, Universiti Kebangsaan Malaysia,43600 Bangi, Selangor, Malaysia

(Received 13 June 2002; accepted 24 March 2003)

Abstract – A new Middle Permian locality in northern Johore, Peninsular Malaysia, yields a small-sized, but compositionally unique, brachiopod fauna consisting of eight species: Pseudoleptodus sp.,Caricula cf. salebrosa Grant, Neochonetes (Nongtaia) aff. arabicus (Hudson & Sudbury), Karavankinasp., Transennatia cf. insculpta (Grant), Hustedia sp., Orthothetina sp., and martiniid indet. The firstfour genera are new records for Malaysia; in particular, the rare taxa Pseudoleptodus and Cariculacharacterize the fauna. The brachiopods occur together with the ammonoid Agathiceras sp., thenautiloid Foordiceras? sp., bivalves, and crinoid stems. The locality belongs to the East Malaya terraneof the Cathaysian biotic region, but some affinities to species of the Sibumasu province are recognized.The Malaysian forms of Pseudoleptodus, Caricula and Transennatia are similar to those of the RatburiLimestone (southern Thailand). A Roadian–early Wordian age is interpreted for the Johore fauna. Thesimilarity of brachiopods reported here with those from the Ratburi Limestone suggests that therewas species interchange or one-way migration between shallow waters of East Malaya and Sibumasuacross the main Palaeo-Tethys. The Tethyan seaway between the two terranes must have been narrowerthan previously interpreted by some authors to allow such faunal traffic during the Roadian–Wordiantime period.

Keywords: Permian, Malaysia, brachiopods, palaeobiogeography.

1. Introduction

The Permian geology of Johore is poorly understood,and Permian marine fossils are known only from a fewrecords, in particular reports of the Geological Surveyof Malaysia (GSM), as compiled by Ibrahim (1987).Igo, Rajah & Kobayashi (1979) described a diverse fu-sulinid fauna (27 species) of probable Kungurian (lateEarly Permian) age from the Sumalayang limestone,southern Johore. The fauna includes two species of theso-called anti-tropical genus Monodiexodina Sosnina,namely M. shiptoni (Dunbar) and M. kattaensis(Schwager). This was the only systematically studiedPermian marine fossil assemblage of Johore.

This paper reports and describes a small faunalassemblage of brachiopods and cephalopods froma new Middle Permian (Guadalupian) exposure inSermin, northern Johore. This is the second record ofPermian brachiopods from Johore, the first being thatof Rahman (1986).

The international Permian time-scale and suggestedcorrelation chart (Jin et al. 1997; Wardlaw, 1999) areutilized in this paper. The Permian fossils reportedhere were discovered by Sone and Leman in August,1999, and additional samples were collected in Marchand October, 2000. All described specimens aredeposited at the National University of Malaysia

† Author for correspondence: msone2@pobox.une.edu.au

(Universiti Kebangsaan Malaysia) with a numberprefixed UKM-F.

2. Regional geology

Prior to this study, the current fossil locality was inclu-ded in the Middle–Late Triassic Gemas (=Semantan)Formation (see the most recent official geological mapof Peninsular Malaysia, Geological Survey of Malaysia,1985). These Triassic clastic sediments are the mostcommon rock type in the central to southern part of theCentral Belt, and are distributed extensively in the westof the present region.

Bachik (1985) first reported the occurrence of marinefossils from the present area, which he considered tobe Triassic in age. Rahman (1986) subsequently listednames of brachiopods and one ammonoid Agathicerasfrom this fauna. The brachiopods were identified asSpiriferellina cf. adunctata Waterhouse & Piyasin,Leptodus sp., Retimarginifera sp., echinoconchid cf.‘Echinoconchus’ fasciatus (Kutorga), and linopro-ductid gen. and sp. indet., and a general Middle Permianage was suggested for this assemblage (C. H. C.Brunton, GSM unpub. report, 1986).

Bachik’s outcrop was a road-cut on a dirt road andwas near a tributary of the Sungai (River) KapehKubang (R. Bachik, GSM, pers. comm. August, 2001).According to the grid reference WL416934 (seeIbrahim, 1987), it should be a few hundred metres

524 M. SONE, I . METCALFE & M. S. LEMAN

Figure 1. Map of the Middle Permian fossil locality in Sermin, northern Johore.

east of the current road-cut outcrop beyond Route 12(Fig. 1). The present area has undergone extensivedevelopment of oil palm plantations and subsequentsevere erosion over the last ten years. The presentauthors failed to relocate Bachik’s locality in the field,and it is uncertain whether it was stratigraphicallyequivalent to the newly found fossil horizon. No otherPermian sediment is known to be distributed aroundthis region.

3. Fossil locality

The present fossil outcrop (02◦ 39′ 06′′ N, 102◦ 51′

51′′ E, by Magellan GPS Tracker) is located in theSermin area on the Kuantan–Segamat Highway (Route12), and is about 2.3 km NE of the T-junction to theagricultural township Felda Pemanis (Fig. 1). It is alarge road-cut on the northern side of the road, and is ap-proximately 20 m in height and 400 m in lateral length,but is mostly covered with vegetation. The fossils werefound in the middle part of the road-cut. The exposureis divided by a normal fault into two lithologicallydistinct sedimentary units. The sequence in whichthe fossils were found is in the foot wall (Fig. 2),and it dips 54◦ W with a strike of N 48◦ W. Anothersequence in the hanging wall dips 70◦ W with a strike ofN 40◦ W.

The fossils were recovered from a band about 8 mbelow the upper boundary of the foot-wall sequence(Fig. 3). The fossil band is about 2 m thick andconsists of brown to purple, tuffaceous, coarse-grained

sandstone and black siltstone. Small fragments ofcrinoid stems are abundant. Brachiopods, cephalopods(ammonoids and one nautiloid) and bivalves are rare.The fossil band is underlain by beds of black shale,sandstone and siltstone. Mud clasts in the lower partof a sandstone layer scraped off from an underlyingshale bed indicate the stratigraphic younging direction(Figs 2, 3). On the other hand, another unit of thehanging wall is made up of massive to thickly bedded,coarse yellowish sandstone. No age-indicative fossilwas found in this sediment; hence the stratigraphicrelationship of the two fault-detached units is unknown.

4. Brachiopod systematics (M. Sone)

Class STROPHOMENATA Williams, Carlson,Brunton, Holmer & Popov, 1996

Order PRODUCTIDA Sarytcheva &Sokolskaya, 1959

Suborder LYTTONIIDINA Williams,Harper & Grant, 2000

Superfamily LYTTONIOIDEA Waagen, 1883Family LYTTONIIDAE Waagen, 1883

Subfamily POIKILOSAKINAE Williams, 1953Genus Pseudoleptodus Stehli, 1956

Pseudoleptodus sp.Figure 4a–i

1976 Pseudoleptodus? sp. indet. Grant, p. 160,pl. 43, figs 1–7.

Permian palaeobiogeography of Malaysia 525

Figure 2. Sketch of the road-cut fossil outcrop studied in this paper. Line A–B corresponds to that of Figure 3.

Figure 3. Stratigraphic column of the fossil sequence.

Size ranges of available specimens. Eight ventral in-ternal moulds were available, of which seven specimensUKM-F441–447 are illustrated, ranging in size from2.5 to 13.5 mm in half width and from 3 to at least18 mm in length.

Description. The assemblage of small shells exhibitsontogenetic stages. In profile, juveniles are cup-shapedand mature specimens are more spatulate. The outlineis ovate. Very fine tubercles are present over the

ventral internal surface of both inlobes and outlobes.Internally, the lobar apparatus varies considerablyin form from shell to shell. In juvenile stages, thevallum (a wall bounding outlobes and inlobes) isvery weak and instead lobar channels are marked bytwo parallel grooves separated by a narrowly roundedmedian thickening (UKM-F441,442; Fig. 4a–c). Inlater but still immature growth stages, the vallumbecomes higher with wide outlobes, as the loops of thevallum are often highly rounded, showing a latilobatecondition (UKM-F443–445; Fig. 4d–g). Lobes becomeangustilobate in later growth stages (UKM-F446,447;Fig. 4h, i). Mature lobation is fairly regular and as manyas six relatively long lateral outlobes are developed.As is common for the genus, outlobe channels arecommonly wider than inlobes.

Remarks. An asymmetrical muscle apparatus is themost obvious characteristic of Poikilosakinae but isnot preserved in the present material. The matureshell UKM-F447 displays extended lateral lobes,atypical for Pseudoleptodus. Hence, there is somepossibility that the Malaysian material may be alliedto the larger Lyttoniinae genus, Eolyttonia Frederiks orCollemataria Cooper & Grant, whose immature formsalso have latilobate to angustilobate features similarto Pseudoleptodus. However, the overall appearanceof the small Malaysian shells is comparable to that ofPseudoleptodus. The present form may represent anadvanced species of the genus.

A Ratburi form of Pseudoleptodus? sp. indet. de-scribed by Grant (1976, p. 160, pl. 43, figs 1–7) isrepresented by three shells. The semi-mature ventralshell in his figures 1–3 is about 19 mm wide andhas angustilobate lobes alike to the Malaysian forms.The mature larger ventral shell in his figure 7 is

526 M. SONE, I . METCALFE & M. S. LEMAN

Figure 4. Pseudoleptodus sp., all specimens are ventral internal moulds. (a) UKM-F441, juvenile shell showing an irregular,asymmetrical lobation and open trough-like median inlobe. (b, c) UKM-F442, left half of a cup-shaped juvenile shell with a weakvallum, in ventral (b) and anterior (c) views. (d) UKM-F443, fragment of inlobate left lateral lobes. (e) UKM-F444, fragment ofinlobate right lateral lobes with rounded loops. (f, g) UKM-F445, shell in ventral (f ) and anterior (g) views; anterior part of a shell(right half ) showing latilobate condition with highly rounded loops of an incomplete vallum. (h) UKM-F446, left half of a possiblemature shell. (i) UKM-F447, right half of a possible mature shell showing angustilobate lobes, strong vallum, and part of hollowmedian inlobe. Scale bar = 10 mm.

20 mm in half width, and displays strongly slantingsolidiseptate lobes. This feature corresponds to theobservation of Cooper & Grant (1974, p. 392) that,in some species of Pseudoleptodus, large and matureshells tend to develop moderately to strongly obliquelateral lobes. Grant’s classification of his species intoPseudoleptodus is supported here. The Thai speciesdevelops five or more lateral outlobes. It is mostcomparable to the Malaysian shells among all knownspecies, although the Thai material shows slightly moreoblique lateral lobes.

The type species of the genus, Pseudoleptodusgetawayensis Stehli (1956, p. 312, pl. 41, figs 2, 5;pl. 42, figs 1, 3; also in Cooper & Grant, 1974, p. 395,pl. 130, figs 18–34), is known from the Getaway Lime-stone Member (upper Roadian–lowermost Wordian)of the lower Cherry Canyon Formation, West Texas.It has relatively regular lobation, with four to fiveoutlobes, and although not the youngest it is the mostadvanced form among all Texan species, approachingthe Southeast Asian representatives. However, theMalaysian and Thai forms have longer lateral lobes.

Cooper & Grant (1974, p. 393) considered someTimorese lyttoniids of Wanner (1935, p. 212, pl. 6,(figs 1, 2 as Cardinocrania), (fig. 8b, c as Poikilosakos))to be Pseudoleptodus. This was noted but not incor-porated into the new Treatise (see Williams, Harper& Grant, 2000, pp. 631, 637). Wanner’s materials,

however, clearly possess asymmetrical ventral musclescars and relatively thick shells, so they are not eitherCardinocrania Waagen (Lyttoniinae, muscle scars onboth sides of the muscle region) or Poikilosakos Watson(thin shell). As for the above-mentioned Timoreseshells, Cardinocrania waageni Wanner (1935, pl. 6,figs 1, 2) probably represents a new poikilosakinidgenus, and the one ventral valve of Wanner (1935, pl.6, fig. 8b, c) included in Poikilosakos variabile Wannerpossibly belongs to Pseudoleptodus in agreement withCooper & Grant (1974).

Waterhouse & Piyasin (1970, p. 134, pl. 24, figs 1–9)described small shells as Leptodus sp. from theRatburi Limestone in Khao Phrik, Thailand. It isoval in outline, with up to six short outlobes. Theysuperficially resemble Pseudoleptodus, but possessa symmetric muscle scar and develop solidiseptateridges, suggesting no relation to Poikilosakinae.

Suborder CHONETIDINA Muir-Wood, 1955Superfamily CHONETOIDEA Bronn, 1862

Family RUGOSOCHONETIDAE Muir-Wood, 1962Subfamily RUGOSOCHONETINAE Muir-Wood, 1962

Genus Neochonetes Muir-Wood, 1962Subgenus Nongtaia Archbold, 1999

Neochonetes (Nongtaia) aff. arabicus

(Hudson & Sudbury, 1959)Figure 5a–l

Permian palaeobiogeography of Malaysia 527

Figure 5. Neochonetes (Nongtaia) aff. arabicus (Hudson & Sudbury, 1959). (a) UKM-F449, internal mould of a juvenile ventral valve.(b) UKM-F450, internal mould of a juvenile ventral valve. (c, d) UKM-F451a,b, dorsal external mould (c), dorsal interior(d) of the same valve (rubber cast). (e) UKM-F452, dorsal external mould. (f ) UKM-F453, ventral internal mould. (g) UKM-F454,dorsal external mould. (h) UKM-F455, dorsal external mould. (i) UKM-F456, ventral internal mould. (j, k) UKM-F457a,b, internalmould of a ventral valve (j), incomplete exterior (k) of the same valve (rubber cast). (l) UKM-F458, ventral internal mould. Scale bar =10 mm.

1959 Chonetes arabicus Hudson & Sudbury,p. 26, pl. 3, figs 6–16; pl. 6, figs 14–18.

1990 Neochonetes (Sommeriella) arabicusHudson & Sudbury; Archbold & Burrett,p. 121, fig. 1A–C.

1999 N. (S.) arabicus Hudson & Sudbury;Angiolini in Angiolini & Bucher, p. 678,fig. 12:15–21.

Size ranges of available specimens. Maximum width4.5–14 mm in range; the maximum width of matureshells is slightly anterior of the hinge. Ventral length3–8.5 mm in range; dorsal length up to 8 mm.

Remarks. This species is a small Neochonetes withrelatively fine capillae. Ventral valves are relativelyconvex. The ventral sulcus is broad and very weaklydeveloped, and the dorsal fold is obsolescent or almostabsent.

Archbold (1999) proposed two new subgenera ofNeochonetes, namely Nongtaia and Zechiella. Theformer is characterized most importantly by its smallsize, while the latter has weak ornament and anobsolescent sulcus as represented by those known inthe Late Permian Zechstein Basin of Europe. The

type species of the former subgenus, Neochonetes(Nongtaia) taoni Archbold (1999, p. 76, fig. 3A–O), is asmall Neochonetes characterized by a narrow, distinctventral sulcus and dorsal fold, and relatively coarsecapillae. It is unlike the present Malaysian form.

Archbold (1999, p. 76) tentatively included Chonetesarabicus Hudson & Sudbury (1959) in Nongtaia. Thisis a tiny species of Neochonetes from Member 1 of thelower Khuff Formation (Wordian) in Oman, previouslyreferred to another subgenus Sommeriella by Archbold& Burrett (1990) and by Angiolini & Bucher (1999).Unlike N. (Nongtaia) taoni, the Arabian species hasrelatively fine capillae and a weakly developed sulcusand fold, and hence is atypical of N. (Nongtaia).The subsurface material of Hudson & Sudbury(1959, pl. 6, figs 14–18) and a recent collection ofAngiolini & Bucher (1999, fig. 12:15–21) exhibitobsolescent folds and sulci, and are comparable tothe Malaysian form. The Malaysian species is slightlylarger than the recorded specimens of N. (Nongtaia)arabicus.

The present chonetids also share some similaritiesto Chonetes variolata var. baroghilensis Reed (1925,p. 40, pl. 3, figs 1–4) from Baroghil, northern Pakistan,

528 M. SONE, I . METCALFE & M. S. LEMAN

Figure 6. Caricula cf. salebrosa Grant, 1976. UKM-F459,dorsal external mould. Shell in ventral (a) and anterior (b) views.Scale bar = 10 mm.

particularly in having strong convexity and a broadsulcus. The appearance of the ventral interior is alsosimilar. The same species Neochonetes (Sommeri-ella) baroghilensis was revised from the Bolorian–Kubergandian (Kungurian–Roadian) beds of Baroghiland Lashkargaz by Angiolini (1996, p. 10, pl. 1,figs 12–17). The Pakistani species, however, differsfrom the Malaysian form in having much largershells (11.5–27.6 mm wide and 7–14.2 mm long)which suggests it belongs to Sommeriella. Likewise,superficial similarities between N. (N.) arabicus andN. (S.) baroghilensis have been noted by Hudson &Sudbury (1959, p. 28) and by Angiolini & Bucher(1999, p. 678), who both, however, differentiated thetwo species by means of their sizes. The Malaysianspecies is here placed provisionally under N. (Nongtaia)in view of its small size and notable affinities to N. (N.)arabicus.

Archbold (1981a, p. 113) pointed out that Kungurianand younger Permian representatives of Neochonetesare small in size. This view was reflected in his newclassification of the four subgenera (see Archbold,1999). It seems possible that the larger N. (S.)baroghilensis had direct evolutionary linkage to eitherN. (N.) arabicus or the present species through theKungurian–Wordian lineage of Neochonetes in thesouthern Tethys.

Suborder PRODUCTIDINA Waagen, 1883Superfamily PRODUCTOIDEA Gray, 1840

Family PRODUCTELLIDAE Schuchert, 1929Subfamily MARGINIFERINAE Stehli, 1954

Tribe Paucispiniferini Muir-Wood & Cooper, 1960Genus Caricula Grant, 1976

Caricula cf. salebrosa Grant, 1976Figure 6a, b

1976 Caricula salebrosa Grant, p. 131, pl. 31,figs 1–30.

Remarks. A single mould of the dorsal exterior UKM-F459 was available, 9 mm wide and 7 mm long. Thissmall dorsal valve is fairly concave in profile and trans-verse in outline, with ears slightly extended. The vis-ceral disc is ornamented strongly by regular rugae butis very weakly reticulated by low costae. Only the trailis well costate. The fold is high anteriorly, and no dorsalspine is present. In all visible morphological characters,the present material is unequivocally assigned to thedistinctive genus Caricula.

The type species Caricula salebrosa Grant (1976)from the Ratburi Limestone in Ko Muk NE is the onlyknown species of the genus. The exterior of theMalaysian dorsal valve appears almost identical to thatof C. salebrosa, of which six specimens for the dorsalexterior were illustrated by Grant (1976, pl. 31, figs 2,3, 11, 15–18, 23, 24, 28, 29). The Thai and Malaysianforms are probably conspecific, but the identificationis tentative due to a lack of ventral information in thepresent collection. Waterhouse, Pitakpaivan & Mantajit(1981, p. 80) also reported Caricula sp. from Ko YaoNoi, peninsular Thailand (of possible Late Artinskian(Baigendzhinian) age according to Archbold (1999,p. 74)), but the material was not illustrated.

Genus Transennatia Waterhouse, 1975

Transennatia cf. insculpta (Grant, 1976)Figures 7a–i, 8c

1976 Gratiosina insculpta Grant, p. 135, pl. 32,figs 1–37; pl. 33, figs 1–16.

Size ranges of available specimens. Maximum hingewidth up to 16 mm (ventral); 9–14 mm (dorsal). Length10.5–15.5 mm (ventral); 7–9 mm (dorsal).

Description. Its small ears and distinctive reticulationare indicative of Transennatia. The shells are medium-sized for the genus. It is subquadrate or slightlytransverse in outline and is weak to moderatelygeniculate in ventral profile. The visceral disc is sharplyreticulated by fine costae and rugae; costae convergeonto the fold. The trail is relatively short. In the dorsalinterior, a median septum is short and low, and anterioradductor muscle pads are non-dendritic and slightlyopened anteriorly.

Remarks. This species is most abundant in the presentfauna. The present form shares some close similaritiesto the Ratburi species Transennatia insculpta (Grant,1976) from Ko Muk; most notably in having arelatively small size, subquadrate outline, feeble ventralgeniculation, very fine reticulation, and opened anterioradductor scars. The Malaysian form is distinguishedfrom the type species Transennatia gratiosa (Waagen,1884, p. 691, pl. 72, figs 3–7) from the Wargal andChhidru formations of the Salt Range, Pakistan, bybeing only about half the size. T. insculpta is also halfas large as T. gratiosa, as compared by Grant (1976,p. 136). Most notably, anterior adductor muscle pads of

Permian palaeobiogeography of Malaysia 529

Figure 7. Transennatia cf. insculpta (Grant, 1976). (a) UKM-F460, posterior part of a ventral internal mould. (b) UKM-F461,incomplete external mould of a dorsal shell. (c) UKM-F462, incomplete internal mould of a ventral shell. (d) UKM-F463, externalmould of a semi-mature dorsal shell; ears probably lost. (e) UKM-F464, dorsal interior of an immature shell (rubber cast). (f ) UKM-F465, dorsal interior (rubber cast) of a semi-mature shell showing anteriorly opened muscle pads. (g) UKM-F466, fragment of aventral? internal mould. (h) UKM-F467, external mould of a semi-mature dorsal shell having relatively coarse ornament. (i) UKM-F468, internal mould of a large mature ventral shell. Scale bar = 10 mm for all, except 12 mm for (i).

Figure 8. Comparison of internal structures of three Transennatia species. (a, b) T. insculpta (Grant) from the Ratburi Limestone of KoMuk, southern Thailand (from Grant, 1976, pl. 33, figs 9, 14). (c) T . cf. insculpta (Grant) from Johore, UKM-F465. (d) Type speciesT. gratiosa (Waagen) from the Kalabagh Member of the Wargal Limestone in the Salt Range, Pakistan (from Grant, 1976, pl. 33,fig. 23). Figure 8a, b, d reproduced with permission of SEPM (Society for Sedimentary Geology).

both the present form and T. insculpta are alike, andare more open anteriorly than those of T. gratiosa asshown by Reed (1944, pl. 19, figs 6, 7) and Grant (1976,pl. 33) (see comparison in Fig. 8).

In addition to the Ratburi T. insculpta, one conjoinedshell specimen from Ko Muk NW (Grant, 1976,pl. 32, figs 15–17) as noted ‘small but apparentlymature shell’ in the caption is more strongly sulcate

and more sturdily costate than those of Ko Muk NE. Asimilar intraspecific variation can also be recognized inthe Malaysian collection as seen in Specimen UKM-F467 (Fig. 7h).

Another Malaysian species Transennatia termi-erorum Sone (in Sone, Leman & Shi, 2001, p. 12,fig. 7:1–10, 13–15) from the early Capitanian of Pahangis characterized by large extended ears and strong

530 M. SONE, I . METCALFE & M. S. LEMAN

Figure 9. (a–e) Karavankina sp., (a, b) UKM-F471, dorsal external mould. (c) UKM-F472, dorsal external mould. (d) UKM-F473,incomplete ventral internal mould (left flank). (e) UKM-F474, incomplete dorsal interior (rubber cast). Scale bar = 10 mm, except5 mm for (b).

geniculation, and its reticulation is coarser than that ofthe Johore forms. Among many other known speciesof the genus, Early–early Middle Permian represent-atives are rather rare. The Kubergandian (Roadian)species Transennatia reedi Angiolini (1995, p. 205,fig. 16:4; also in Angiolini, 1996, p. 14, pl. 1, figs 27,28; pl. 2, figs 1–4) from northern Pakistan is large andis coarsely reticulated, with costae not converging intoa sulcus.

Retimarginifera praelecta (Reed, 1925) sensuAngiolini (1996, pl. 1, figs 25, 26) from the Bolorian–Kubergandian (Kungurian–Roadian) of northernPakistan has notably sturdy, coarse costae whichconverge into a deep sulcus and diverge on flanks. Itsappearance and description seem to suggest a morereasonable assignment to Transennatia rather thanRetimarginifera. Its ornament is much coarser than theMalaysian form.

Superfamily ECHINOCONCHOIDEA Stehli, 1954Family ECHINOCONCHIDAE Stehli, 1954

Subfamily ECHINOCONCHINAE Stehli, 1954Tribe Karavankinini Ramovs, 1966Genus Karavankina Ramovs, 1966

Comments. The genus Karavankina was fully de-scribed and defined in Ramovs (1969). His 1969 pub-lication, in which he proposed this new generic name(and the new subfamily name Karavankininae and fournew species-group names), was originally intended tobe published in 1966. Prior to its publication in 1969,Ramovs (1966) first published the names accompaniedby fixation of the type species Karavankina typica(as indicated on the figure) and by comparison ofthe genus with other pre-existing taxa. This 1966paper satisfies the provisions of Articles 13.1–3 of theInternational Code of Zoological Nomenclature, and

therefore establishes priority for all the names madeavailable in Ramovs (1966). Karavankina Ramovs,1969 [=Karavankina Ramovs, 1966, nomen nudum]as stated in the new Brachiopoda Treatise (Bruntonet al. 2000, p. 512) is incorrect. Sarytcheva (1968)accepted Karavankina Ramovs, 1966, yet as a subgenusof Echinoconchus Weller.

Karavankina sp.Figure 9a–e

Description. The species is small-sized (10.5–16 mmwide and 7.5–10 mm long in dorsal valves) for thegenus. The outline varies from transverse to subcircu-lar. Coarse, high relief concentric bands consisting ofalternating smooth and spinous regions are indicativeof Karavankina. Numerous spines are present on bothvalves. Ventral external spines are oblique anteriorly.Dorsal external spines are erect and are regularlyarranged as differentiated by size. Fine, weak wrinklesare observed on dorsal spine-free bands transversely.Dorsal endo-spines are present.

Remarks. Many Carbo-Permian species of Karavank-ina share considerable superficial similarities. Exam-ination of dorsal spine patterns may help with moreeffective identification at the species level, althoughthese features are not sufficiently understood in mostknown species. The type species Karavankina typicaRamovs (1966, fig. 7; also in Ramovs, 1969, p. 254,pl. 1, figs 1–4) from the Middle Permian of northeasternSlovenia is similar to the Malaysian shells in overallappearance, but is a little larger.

Another Slovenian echinoconchid has been knownfrom the Karawanken Mountains, originally as Pro-ductus elegans M’Coy sensu Schellwien (1900b,p. 52, pl. 8, figs 14–17). This was later revised to

Permian palaeobiogeography of Malaysia 531

Figure 10. (a–d) Hustedia sp., UKM-F475, conjoined shells in ventral (a), dorsal (b), right lateral (c) and left lateral (d) views.(e) martiniid indet., UKM-F476, ventralinternal mould. (f–j) Orthothetina sp., (f ) UKM-F477, internal mould of an immature dorsalvalve. (g) UKM-F478, incomplete internal mould of a large mature shell’s beak in ventral view. (h–j) UKM-F479a, internal mould ofa semi-mature ventral valve in ventral view (h), interarea in dorsal view (i), (j) UKM-F479b, incomplete exterior (rubber cast) of thesame ventral valve. Scale bar = 5 mm for (a–d), 10 mm for (e, g), 6.7 mm for (f ), 8 mm for (h–j).

Karavankina schellwieni Ramovs (1966, fig. 6a–c;also in Ramovs, 1969, p. 264, pl. 2, figs 1–4) withadditional material. It was also reported from theNeoschwagerina craticulifera fusulinid bed (Wordian)of the Julian Alps, northwestern Slovenia (Flugel,Kochansky-Devide & Ramovs, 1984, pl. 39, fig. 8).K. schellwieni is a relatively small species with narrowdorsal concentric bands, and appears comparable tothe present form, although precise identification isnot possible without dorsal spinous details of theSlovenian species. The true P. elegans M’Coy hasnow been designated as the type species of the mid-Carboniferous genus Echinoconchella Lazarev (1985,p. 70).

Echinoconchus cristatus Reed (1931, p. 12, pl. 2,figs 7–10a), proposed as a variation of E. fasciatus(Kutorga), from the Bamyan limestone (Wordian) ofcentral Afghanistan, is interpreted as a species ofKaravankina, based on its characteristic concentricbands. It is about twice as large as the Malaysian form.E. fasciatus has now been placed under Karavankina(see Sarytcheva, 1968, p. 95).

Laotian (Kham-keut) and Vietnamese (Van-yen)echinoconchids both from the Pseudoschwagerinaprinceps fusulinid beds (Early Permian) were illus-trated as Productus elegans by Mansuy (1913, p. 30,pl. 2, fig. 3a–d). They also unequivocally belong toKaravankina owing to their possession of high-reliefbands. Their dorsal views are not available, preventingdetailed comparison to other species.

The Malaysian form exhibits a distinctive dorsalspine pattern (Fig. 9b), and thus may represent a newspecies. However, more morphological information,particularly on the ventral valve, is required before anew species can be established.

Order ORTHOTETIDA Waagen, 1884Suborder ORTHOTETIDINA Waagen, 1884Superfamily ORTHOTETOIDEA Waagen, 1884

Family MEEKELLIDAE Stehli, 1954Subfamily MEEKELLINAE Stehli, 1954

Genus Orthothetina Schellwien, 1900a

Orthothetina sp.Figure 10f–j

Remarks. Generic assignment to Orthothetina is veri-fied most precisely by a non-plicate orthotetidine shellwith a pair of sub-parallel dental plates extending one-third of the valve length. It is medium-sized for thegenus, with the greatest width at mid-valve, and issemi-ovate in outline, with a short beak. Surface isornamented by radial capillae of uneven width.

In Pahang (Peninsular Malaysia), two other formsof Orthothetina have been found: Orthothetina sp.in the Jengka Pass shale of the late? Middle Permian(Nakazawa, 1973) and O. cf. iljinae Sokolskaya inthe early Capitanian of the Bera Formation (Sone,Leman & Shi, 2001). They are not similar to the Johorespecies.

532 M. SONE, I . METCALFE & M. S. LEMAN

Class RHYNCHONELLATA Williams, Carlson,Brunton, Holmer & Popov, 1996Order ATHYRIDIDA Boucot,

Johnson & Staton, 1964Suborder RETZIIDINA Boucot,

Johnson & Staton, 1964Superfamily RETZIOIDEA Waagen, 1883

Family NEORETZIIDAE Dagis, 1972Subfamily HUSTEDIINAE Grunt, 1986Genus Hustedia Hall & Clarke, 1893

Hustedia sp.Figure 10a–d

Remarks. A single specimen UKM-F475 of slightlydeformed conjoined shells was available. It is 6 mmwide and 8 mm long. Two Ratburi species, namelyHustedia ratburiensis Waterhouse & Piyasin (1970)(=H. nakornsrii Yanagida, 1971 (imprint 1970)) andH. funaria Grant (1976), are known. The Johore formis more similar to the former than the latter whichis distinctively long and narrow in outline. Furthercomparison is difficult due to the limited material.

Order SPIRIFERIDA Waagen, 1883Suborder SPIRIFERIDINA Waagen, 1883Superfamily MARTINIOIDEA Waagen, 1883

Family MARTINIIDAE Waagen, 1883

martiniid indet.Figure 10e

Remarks. One ventral internal mould UKM-F476reveals straight pallial markings, a muscle field bisectedby a median groove, and smooth lateral slopes,suggestive of Martiniidae. Generic assignment is notpossible because of the limited collection.

5. Correlation and age

Two species of cephalopods and eight species of bra-chiopods are recognized in the present fauna (Table 1).The ammonoid Agathiceras sp. (Fig. 11a) indicates apre-Capitanian age for the fauna. The genus rangesfrom the Moscovian (Late Carboniferous) through tothe Wordian, with greatest abundance in Roadian andWordian rocks (Glenister et al. 1990; Zhou et al.1999). Two other Malaysian forms, both as Agathicerassp., were described from two horizons of the BeraFormation in Pahang: one from a Roadian or Wordianbed of Felda Mayam (Sone & Leman, 2000) and theother from the Wordian cephalopod fauna of BeraSouth (Sone, Leman & Ehiro, 2001). The Felda Mayamform has a flattened lateral flank and high umbilicalangle, resembling the Johore shells. The nautiloidFoordiceras? sp. (Fig. 11b) is also discriminated; it haslittle stratigraphic value.

The key brachiopod species of Johore belongs tothe extremely rare genus Caricula, as it is probably

Table 1. Middle Permian cephalopods and brachiopods of Sermin,northern Johore, Peninsular Malaysia; cephalopod shellsundifferentiated

Number of samples

Species Ventral Dorsal Conjoined

CephalopodsAgathiceras sp. 4Foordiceras? sp. 1BrachiopodsCaricula cf. salebrosa 1

Grant, 1976Pseudoleptodus sp. 7Transennatia cf. insculpta 4 9

(Grant, 1976)Neochonetes (Nongtaia) aff. 6 4

arabicus (Hudson &Sudbury, 1959)

Karavankina sp. 1 3Hustedia sp. 1Orthothetina sp. 2 1martiniid indet. 1

Figure 11. Two mid-Permian cephalopods from northern Johore.(a) ammonoid Agathiceras sp., UKM-F480. (b) nautiloidFoordiceras? sp., UKM-F481. Scale bar = 10 mm for (a),7.5 mm for (b).

conspecific to C. salebrosa Grant of Ko Muk NE,which is the only known species of the genus. Thissignifies linkage to the Ratburi fauna. The occurrenceof Pseudoleptodus sp. is also significant. This genusis rare in the Tethys, having been known only in theRatburi Limestone of Ko Muk NE, although in WestTexas it occurs from the Wolfcampian to throughout theGuadalupian (Early–Middle Permian). The Malaysianform appears to be a very advanced species. Third,the Johore form of Transennatia again resembles theRatburi T. insculpta (Grant) of Ko Muk NE and NW.Transennatia is common through the Middle to LatePermian, but is extremely rare or absent in the EarlyPermian. Thus, the similarities of Johore forms ofCaricula, Pseudoleptodus and Transennatia to the Thaispecies strongly suggest correlation to the Ko Muk (NEand NW) horizons of the Ratburi Limestone.

The genus Orthothetina is most common duringthe Guadalupian–Lopingian (Middle–Late Permian).The genus Karavankina ranges in age from the LateCarboniferous to the Wordian, except the Oman form ofYanagida & Pillevuit (1994) whose age may be slightly

Permian palaeobiogeography of Malaysia 533

younger suggested by foraminifers (see Angiolini &Bucher, 1999, p. 674). The two genera are not knownfrom the Ratburi Limestone.

The age of the Ratburi Limestone brachiopodshas been of great dispute. Age assignments havediffered from author to author, for instance, Kazanian(Waterhouse & Piyasin, 1970), late Artinskian(Yanagida, 1970; Grant, 1976), Kungurian(Waterhouse, 1973, 1981), late Artinskian–earlyKungurian (Shi & Archbold, 1995), and Roadian–Wordian (Angiolini & Bucher, 1999). Archbold (1999)suggested an Ufimian age (roughly equivalent to theRoadian of the international standard, see Archbold,1998), based primarily on faunal similarities withBitauni Timor, West Irian Jaya and Western Australia.Comprehensive biostratigraphic analyses of theRatburi Limestone were provided by Baird, Dawson &Vachard (1993) and Fontaine et al. (1994a).

The Ratburi brachiopods have been considered to beto some degree similar to those of the Amb fauna ofPakistan (Yanagida, 1970; Grant, 1976) and the Khufffauna of Oman (Angiolini et al. 1998; Angiolini &Bucher, 1999). The similarities are largely in thegeneric level. Based on studies of Gomankov & Burov(1999), Iqbal et al. (1998) and Wardlaw & Pogue(1995), the age of the Amb Formation can be con-strained to a range of Bolorian–Kazanian (Kungurian–Wordian). Angiolini (2001a, p. 312) considered theAmb brachiopod fauna to be no younger than mid-Wordian, based on brachiopod correlations. Yet, arecent conodont study shows that an even slightlyyounger age is possible for the upper part of the AmbFormation (see Mei & Henderson, 2001; Wardlaw &Mei, 1999). The Johore brachiopods reveal no definablelinkage to the Amb fauna.

On the other hand, the Johore Neochonetes sharesnotable similarities with the Oman N. (Nongtaia)arabicus (Hudson & Sudbury), which suggests possiblecorrelation with Member 1 of the lower KhuffFormation. The Khuff fauna was assigned by Angioliniet al. (1998, 2003) to a Wordian age, which seems mostlikely, considering all fossil data.

Kotlyar et al. (1999) reported a brachiopod as-semblage from limestone blocks of the Marta RiverBasin, Crimea, which curiously includes severalRatburi species, although it is yet undescribed. TheCrimean brachiopods are associated with many fu-sulinids including index fossils Neoshwagerina simplexOzawa and Presumatrina neoshwagerinoides (Deprat)and with Wordian ammonoids such as Tauroceras wan-neri Toumanskaya. Kotlyar et al. (1999) assigned thisfauna to a late Kubergandian age. Leven (1998), how-ever, assigned the N. simplex–P. neoshwagerinoideszone to the lower Murghabian. Jin et al. (1997) andJin & Shang (2000) correlated the same biozone to themiddle–upper Roadian. A late Roadian/early Wordianage for the Crimean fauna seems most probable in viewof fusulinid and ammonoid biozonations.

On balance, as well as the Ufimian suggested byArchbold (1999), the early Wordian is here consideredalso possible for the age of the Ratburi brachiopodfaunas. A Roadian–early Wordian age is favouredfor the Johore fauna. This is based on correlation tothe Ratburi brachiopod horizons, the presence of theadvanced form of Pseudoleptodus, and the specificaffinity of the small Wordian species Neochonetes(Nongtaia) arabicus.

6. Palaeobiogeographic implications

The East Malaya terrane, on which the present fauna islocated, has been considered to be an extended part ofthe Indochina block and has been broadly referred to asa unit of the tropical Cathaysian region throughout thePermian (e.g. Metcalfe, 1998, 2002; Shi & Archbold,1998). The present Johore brachiopods, however, showsome faunal linkage to the Sibumasu province (Fang,1991) of the Cimmerian region.

The Johore assemblage as a whole represents awarm-water Tethyan-type fauna, which, however, con-tains no diagnostic Cathaysian (South China) generaand species. Of seven genera identified, Cariculaand Pseudoleptodus can be referred to as Sibumasuelements, as they are known, elsewhere in the Tethys,only in peninsular Thailand. The other four, Transenna-tia, Karavankina, Orthothetina and Hustedia, suggesta general warm-water environment, as they weredistributed throughout shallow waters of the Tethys,although not restricted to the Cathaysian region; thefirst three are endemic to the Tethyan ocean.

The presence of a mid-Permian Karavankina inJohore implies another non-Cathaysian element in EastMalaya/Indochina. This echinoconchid genus is quitecommon in Carboniferous rocks of South China. Itseems, however, to have disappeared from South Chinasince the Early Permian, while it still persisted in someother regions of the Tethys, including Indochina and theCimmerian region, up to the Middle Permian. (Note:Echinoconchus mapingensis Grabau (1936) from theso-called ‘Maping Limestone brachiopod fauna’ ofGuizhou, South China, is a species of Karavankina(or Echinoconchella), but the age is most likelyearly Bashkirian (mid-Carboniferous) (see Liao, 1999,p. 319)).

At the species level, the Johore forms of Caricula,Pseudoleptodus and Transennatia are more or lesssimilar to those of the Ko Muk limestone, suggestingspecific linkage between East Malaya and Sibumasuover the main Palaeo-Tethys ocean (Fig. 12). AnotherJohore form compared to Neochonetes (Nongtaia)arabicus of Oman may also suggest linkage tothe Cimmerian biota, with particular respect to theemended Sibumasu province of Angiolini (2001b)which includes the Roadian–Wordian of Oman and theSalt Range.

534 M. SONE, I . METCALFE & M. S. LEMAN

Figure 12. Southeast Asian map showing the major tectonicterranes, the main Palaeo-Tethys Suture and other suture zones,the locations of Sermin on the East Malaya terrane and KoMuk on the Sibumasu terrane, and the area with discontinuousPermian Ratburi Limestone (after Fontaine et al. 1994a). Boththe Chiang Mai suture zone and the Simao terrane modified afterMetcalfe (2002).

Possible influences of Sibumasu faunas or sub-tro-pical climates are also tangible in Permian brachiopodsof the Indochina block. Hogeboom & Archbold (1999,p. 260) noted that some Early and Late Permian formsof Laos are referable to Stictozoster Grant, a genuscommon to the Sibumasu and Westralian provinces.Termier & Termier (1970, p. 451, pl. 30, figs 3,10) described Megousia sp. from the mid-PermianSisophon Limestone of Cambodia, which (if not aspecies of Anidanthus Whitehouse) may also suggestan anti-tropical climatic influence, distinctive fromSouth China.

Such faunal linkage between Sibumasu and EastMalaya/Indochina is also recognized in distributionsof two Midian foraminifers, Hemigordiopsis Reichel

and Sphairionia Nguyen (the latter may be incer-tae sedis). The two genera are found in both theEast Malaya/Indochina and Sibumasu terranes, andthroughout the Cimmerian region, but are absent inthe South China block (see Fontaine et al. 1994a,b;Fontaine, Suteethorn & Vachard, 1998; Pronina, 1996;Nestell & Pronina, 1997; Vachard et al. 2001, andreferences therein).

A similar and biogeographically even more res-tricted linkage is indicated by the distribution of lateMurghabian–Midian fusulinid Pseudofusulina padan-gensis (Lange). The species is restricted to Sibumasu(the Ratburi Limestone), East Malaya (Padang in WestSumatra and Jengka Pass in Malaysia) and Indochinaterranes (see Fontaine et al. 1994b; Ueno, Sugiyama& Nagai, 1996), and occurs in both sides of the mainPalaeo-Tethys.

The distribution pattern of the Wordian ammonoidTauroceras scrobiculatum Toumanskaya and its alliesis shown to extend from the western Tethys throughthe Cimmerian region up to East Malaya (Sone, Leman& Ehiro, 2001). They are, however, not known fromIndochina or South China of the main Cathaysian re-gion. Monodiexodina in southern Johore (Igo, Rajah &Kobayashi, 1979) suggests a possible Sibumasu bioticinfluence on East Malaya in the late Early Permian.

Thus, the Permian marine biota of East Malaya/Indochina is not exclusively of the tropical Cathaysiantype, but includes some minor Sibumasu/Cimmerianelements, likely indicating sub-tropical climatic epis-odes. The Bentong-Raub suture zone of PeninsularMalaysia is widely accepted as representing the mainPalaeo-Tethys in Southeast Asia and the palaeobi-ogeographic boundary between the Gondwanan andCathaysian floral provinces (Metcalfe, 2000). This is ingood agreement with general distributions of faunaland floral groups. However, as outlined above, theGondwana–Cathaysia divide between Sibumasu andEast Malaya appears not to have precluded some faunalexchange during the mid-Permian.

It is perhaps too early to discuss the Permian palaeo-geographic relationship between East Malaya andIndochina at this stage, because of the limited dataavailable. Sone, Leman & Shi (2001) concluded thatearly Capitanian brachiopods of the Bera Formationin East Malaya have linkage to the correlative fauna ofIndochina (western Cambodia) but have weaker affinityto those of South China, although the Malaysian faunais broadly referable to the Cathaysian type. On theother hand, the older Johore fauna reported here showsa weaker (if any) connection to Indochina and SouthChina than the Capitanian Bera fauna. Further work isrequired to reveal detailed bio-provincial characters ofEast Malaya and Indochina in comparison to the typicalCathaysian marine biota of South China.

Archbold (1981b, 1991), Archbold & Shi (1995)and Archbold et al. (1982) suggested that late EarlyPermian brachiopod faunas of Bitauni Timor and

Permian palaeobiogeography of Malaysia 535

West Irian Jaya also share close similarities to thoseof the Ratburi Limestone. A similar conclusion wasmade by Sakagami (2000) in terms of bryozoanpalaeobiogeographic affinity. All those brachiopodfaunas, however, contain more or less Gondwanan(Westralian) affinity taxa, which are not associated withthe Johore fauna. This suggests that East Malaya was inwarmer climates and/or in lower latitudes than Timor,West Irian Jaya and Sibumasu of eastern Cimmeria.

In conclusion, there was specific interchange orone-way migration of some brachiopods between theshallow waters of the East Malaya and Sibumasuterranes across the main Palaeo-Tethys during theRoadian–Wordian period. This perhaps suggests a sub-tropical climatic episode for East Malaya at this time.The mid-Permian Palaeo-Tethys seaway between thetwo terranes must have been narrower than previouslyinterpreted to allow such faunal traffic. A vast ocean be-tween Sibumasu and East Malaya/Indochina as recon-structed by Ricou in Baud et al. (1993), Ricou (1995)and Stampfli (2000) appears to be highly unlikely.

Acknowledgements. M. Sone’s Ph.D. study on which thispaper is based is supported by both a University of NewEngland Research Scholarship and an International Post-graduate Research Scholarship. I. Metcalfe acknowledgesongoing support by the Australian Research Council. Wealso acknowledge funding from two IGCP Grants-in-Aid.Critical reviews from Professors N. W. Archbold and A. J.Barber improved this article. Our fieldwork was supportedby the Malaysian Government IRPA Project (02-02-0012-EA186) and the National University of Malaysia FST Project(ST-014-2002). University of New England’s Divisions of′Earth Sciences′ and ′Archaeology and Palaeoanthropology′

provided Sone with laboratory and darkroom facilitiesrespectively. Permission to reproduce photographs from thePalaeontological Society Memoir 9 used in Figure 8a, b, dwas kindly granted by SEPM (Society for SedimentaryGeology).

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